Brugal & Valente 2007 - Dynamic of Large Mammalian Associations in the Pleistocene of Portugal

8/8/2019 Brugal & Valente 2007 - Dynamic of Large Mammalian Associations in the Pleistocene of Portugal

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From the Mediterranean basin to the Portuguese Atlantic shore: Papers in Honor of Anthony Marks Actas do IV Congresso de Arqueologia Peninsular

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Dynamic of large mammalian associations in the Pleistocene of Portugal

Mealhada also presents acheulean artefacts, and a U/Th dateof 329 70 ky is given for Condeixa. Two other sites are situ-

ated in the Almonda karst system: Nascente in the deep karst(with a U/Th date around 150 ky) and Galeria Pesada / Bre-cha das Lascas (or Aroeira).

The straight-tusked elephant Elephas (Paleoloxodon) anti-quus is the most common species (n = 5 occurrences); theyare often old finds, probably found because of the large sizeof bone and teeth of this species. Other Middle Pleistocenespecies are: Hippopotamus incognitus (n = 2) and Homothe-rium latidens (n = 1), the sabertooth felid, as well as taxa stillpresent during the Upper Pleistocene: Cervus elaphus (n = 2),Bos primigenius (n = 1), Equus caballus ssp. indet.(n = 2),Capra pyrenaica (n = 1). Homotherium, determined from oneastragalus, is found in fluviatile sequence of Mealhada given

as Riss deposits (Antunes 1986); this species is presentbetween 900-300 Ky in Western Europe and the Portuguesespecimen could indicate the probable survival of the speciesin this territory3.

One of the most important Middle Pleistocene sites wasfound recently; its Galeria Pesada/Brecha das Lascas loca-ted in Serra da Aire near Almonda, not far from Torres No-vas. A. Marks and his team (Marks et al. , 2002) excavated itbetween 1997-2002 and its thick karstic filling (c. 3 m) hasyielded rich lithic and faunal remains, with an ESR/U/Th datearound 241+ 30-22 Ky (level B2). It is a multilayered sites (sixmain levels in Galeria Pesada) with relatively homogenousremains through the sequence. Globally fifty species are pre-sent from bats, rodents, birds, reptiles, leporids, to mediumand large mammals. Thirteen species of mammals are iden-tified, with the dominance of two cervid species, indicating anopen forest and grassland under a cool climate. One the mostremarkable paleontological contributions of the site concernsthe identification of some new species in the Pleistocene ofPortugal (Brugal, 2004a, and in prep.): archaic Homo (Trin-kaus et al., 2003), Macaca sylvanus (Barbary ape), Damaclactoniana (Clacton deer), Hemitragus sp. (Thar or Asiaticmountain goat), Ovis ammon (wild sheep), Canis gr.etruscus-mosbachensis (archaic wolf), Cuon priscus (Asiatic dhole).They are associated with Cervus elaphus ssp., Equus aff.

mosbachensis, Stephanorhinus hemitoechus , Sus scrofa,Ursus arctos and Castor fiber. Galeria Pesada constitutes themost impressive Middle Pleistocene Portuguese site with ho-minid activities, original lithic industry and rich and originalmammal associations.

UPPER PLEISTOCENE

The data from the first part of this period are scanty andwe can note the persistence ofElephas (Paleoloxodon) anti-quus (isolated finds) in three last interglacial (OIS 5 s.l. )

open-air localities (Casal do Torquato, Carregado, Meirinha).Contrastingly, the period covering the second part of Upper

Pleistocene is the best represented with many sites yieldingfaunal accumulations. They are mostly dated from the LastGlacial, OIS 4 to 2 (c. 70-10 Ky), with some of them from thevery early phases of OIS 1, and are associated with late Mid-dle to late-very late Palaeolithic cultural (n = 21 sites) or natu-rally deposited layers (n = 10). These sites are essentially lo-cated at limestone region of Estremadura in Central Portugalin relatively low elevation and 74% represents cave deposits;few are in cliffs (rockshelter), in pit-falls (algares: Goldra, Cas-cais, Joo Ramos) or in open-air (Quinta do Gaio, Santo An-to do Tojal, Foz do Enxarrique). The number of sites/levelswith mammalian faunas decreases throughout the late upperPleistocene (see Table 1). Most of the archaeological sites are

multileveled either corresponding to the same cultural period(ex. Oliveira) or ranging through several techno-complexes(ex. Lapa do Picareiro, Buraca Escura).

Overall, 57 levels can be considered with reliable faunalidentifications, supported by almost 70 radiometric dates,mainly conventional radiocarbon ones. The dates concernsites not older than 40/35 Ky until around 10 Ky BP. We ar-ranged the dated sites and its levels (uncalibrated BP), plusor minus one sigma, according to the four main techno-com-plexes (Fig. 1): Late Mousterian (MP), Aurignacian-Gravettian(i.e. Early Upper Paleolithic; EUP), ProtoSolutrean-Solutrean(PS + Sol) and Magdalenian (Ma) (adapted from Gamble1986: 135-136). The few ESR and U/Th dates have been ex-cluded of the Fig. 1 in order to keep a more coherent frame-work. We have also indicated the different Heinrich events (H)and Younger Dryas (YD) for this time period, which representmillennium-scale cold events (e.g., Bard, 2002). These abruptevents have clear incidence on the Iberian margin with thepolar front getting to lower latitudes as northern Portugal(42 N during the last glacial maximum, like demonstrated byseveral occurrence of ice-rafted debris [IRD] in cores off thecoast of Portugal), accompanied by drastic cooling of up to4-5 C. Pollen, associated with foraminifer, dinoflagellate cystor18O measurements, documents the impact of these short-term environmental changes on continent (Sanchez-Goni et

al., 2000; Turon et al. , 2003). Six Heinrich events are knownduring the last glacial with dates centred around 14.5 Ky (H1),21-22 ky (H2), 27 Ky (H3), 35 Ky (H4), e.44 Ky (H5) ande.55 Ky (H6), plus the YD of shorter duration (10.5 Ky) (Grou-sset, 2001). Although the accuracy of the radiocarbon agescalibration should be questioned (Fontugne, 2004; and seedErrico and Sanchez Goni, 2003: 771) it seems from the dis-tribution of dated sites (taking one sigma; see Fig. 2) that the

3 New find from the North Sea, dated to ca. 28 Ky, seems to indicate apossible late survival of this species in some parts of Europe (Reumer, etal., 2003).


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different techno-complexes are more or less bracketed inbetween these cold stadials, with relatively continuity and

short overlap (MP-EUP); a clear gap occurred between theSolutrean and Magdalenian cultures. We have then decidedto pool all the different mammal associations into these fourcultural periods in order to summarize all the data and betterappreciate the animal community changes. We can assumethat these repeated cold events have effectively affected themammalian distributions as they did on vegetation composi-tions in Iberia (Sanchez Goni et al., 2002).

Land paleoenvironmental data for the Last Glacial in Por-tugal are extremely limited, with few and dispersed informa-tion obtained from geology, palynology and anthracology, andon faunal record (micro and macro). Different researchershave recently proposed preliminary synthesis on a regional

basis (e.g., Zilho, 1990, 1997; Bicho, 1999; Haws, 2004) andwe can immediately notice that the information is far to becontinuous for this period. A better chrono-climatic and pa-leoenvironmental framework, at least for OIS3, has been ob-tained from deep-sea cores collected off the Iberian margin(dErrico and Sanchez Goni, 2003). A first issue concerns theevidence of a cold peak identified as the Last Glacial Maxi-mum (LGM) not only in the mountain area of northern Portu-gal (periglacial condition as in Serra Estrella, for instance) butalso in the lower areas, such as in Caldeiro cave (basalSolutrean) in Estremadura, based on magnetic susceptibilityfrom cave sediments (Ellwood et al., 1998). During cold anddry phases, the vegetation obtained from pollens and char-coal studies indicates the presence of steppe grasses andherbs as well as forest-parkland wi th coniferous (often domi-nating arboreal spectrum), oaks and birch, which probably ex-panded during interstadial conditions from some refugiumwith Mediterranean trees and shrubs (Quercus , Olea, Pista-cia). These interstadial oscillations were of short duration,known as part of Dansgaard-Oeschger events (D-O), whichregularly allow more or less limited forest expansion. Eleva-tion plays a main role in the distribution of the phytocoenoseas well as in the presence of some mammal species in somesites (especially caprines). It is a complex climatic period witha succession of cold and warm fluctuations (at least about

thirty for OIS3) and the mainland faunal records have a lowerresolution when compared with deep-sea or ice core informa-tion. Deglaciation is also marked by several short climaticphases (D-O) and it is difficult, according to radiocarbon dates(and their error) from the sites, to precise the correlation be-tween cultural levels and climatic peaks. Lastly, during theLGM the sea level was almost 100-120 m below the actualshoreline and in Estremadura the coast was about c. 35 kmeastward (see the dynamic reconstructions of coastline be-tween 18 Ky and present day by Dias, 2004: Fig. 2; also seeour Fig. 1). The change of available land surface, especiallyduring the Solutrean and Magdalenian, probably affected our

archaeological record with disappearance of numerous sites.During the Last Glacial, twenty-three medium to large-

sized terrestrial mammal species are present in the faunalrecord of Portugal: eleven species of Herbivores, eleven spe-cies of carnivores and one large rodent, the b eaver (Table1). The total number of occurrence is related to the numberof levels per period. Among herbivores we can strongly ques-tion the presence of the arctic genus Mammuthus in Portu-gal, only identified by one fragmented tooth lamellae in theMiddle Palaeolithic level of Gruta da Figueira B rava (Antu-nes, 2000). Another remain is a badly preserved shaft of longbone (femur) from Algar de Joo Ramos attributed to thisgenus exclusively on the basis of a younger radiocarbon date(14 170+/-330 BP) and the assumption ofElephas antiquusextinction by this period4. This last species is still known dur-

ing OIS4 from isolated finds at the terrace deposits of SantoAnto do Tojal and several remains found in the Middle Pa-laeolithic level of Foz do Enxarrique dated around 34 Ky byU/Th (e.g., Brugal and Raposo, 1999); it is probably the lastoccurrence of this species in Western Europe (see Stuart,2005). Among cervids, the fallow deer Dama dama is poorlydocumented with one fragmented maxillary with teeth fromMousterian levels of Gruta Nova de Columbeira and only oneisolated tooth from Final Pleistocene of Algar de Joo Ramos(Cardoso, 1989). For carnivores, Cuon alpinus is present withonly one occurrence (fragment of mandible with P4 and M1)from Middle or Upper Palaeolithic levels of Escoural (Cardoso,1992; see also Gautier, 1996 for this cave); this species isnot mentioned in the table due to problems of stratigraphicalposition. We have not reported sea mammals, however theyare present in the Mousterian deposits of Figueira Brava(Antunes, 2000), a cave actually located in a sea cliff: oneremain of ringed seal Pusa hispida and six vertebrae of com-mon dolphin Delphinus delphis, demonstrating colder condi-tion for this period (as the presence of the great auk Pinguinisimpennis ); also one find (small vertebrae) is possibly attri-buted to a cetacean found in the Final Gravettian of LagarVelho (TP06, Moreno-Garcia and Pimenta, 2002).

Globally, for the last glacial the most frequent species arerespectively (by decreasing abundance):

Herbivores: red deer (C. elaphus), horse (E. caballus),aurochs (B. primigenius), and wild goat and boar ( C. py-renaica, S. scrofa);

Carnivores: fox (V. vulpes ), lynx and wolf (L. pardina,C. lupus), brown bear and cave hyena (U. arctos, C.spelaea ).

Some taxa are rare. And besides E. antiquus and Dama,it is noticeable the quasi-absence of the hemion-like horseEquus hydruntinus, occurring only during Middle Paleolithic

4 A better alternative explanation is the possible mixture of faunas of dif-ferent ages, especially in what concerns this type of site, a pit-fall.


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(Pedreira das Salemas) and Solutrean (Caldeiro, Algar deCascais) periods; the latter co-related with the LGM. The pre-

sence of this small equid should imply dryness in climatic en-vironment. The chamois Rupicapra is an uncommon elementalthough present in all considered periods: it is never abun-dant and often is associated to hilly or high elevated sites (ex.Magdalenian of Lapa do Picareiro). Among carnivores, thearchaic species Hyaena prisca was only found in the MiddlePalaeolithic levels of Furninha where an important and well-preserved population is represented. The cave lion was neverabundant with its last occurrence on the Early Upper Palaeo-lithic level of Caldeiro (27-22 000 years BP; Davis, 2002).

Throughout the last glacial sequence, the most outstan-ding process concern the extinction of pachyderm species(proboscid and rhinocerotid) among herbivores and large-

sized predators such hyenids and felids. The taxonomic diver-sity, expressed in number of species per period, shows inte-resting features with a clear herbivore decrease between theLate Mousterian (n = 11 species) and Early Upper Palaeolithic(n = 8), due to the disappearance of pachyderms and fallowdeer, and the scarcity of Equus hydruntinus and Rupicapra .The Upper Palaeolithic is marked by a relative stability in her-bivore structures and then a second decrease of species bythe end of Palaeolithic time with only five or six herbivore spe-cies still existing during the Holocene. Those species are tem-perate elements found in all southern mammal associations(Brugal et al., 2004b, c): Cervus elaphus, Capreolus capreo-lus, Sus scrofa, Capra pyrenaica5, Bos primigenius and Equuscaballus (?); the last two species disappear at the end ofHolocene or earlier.

We observe a different pattern for carnivores with a gra-dual decrease followed by a clear decline just after the LGM,between Solutrean and Magdalenian (see Valente, 2004, andthe example of Caldeiro in Davis, 2002), and then only atthe end of Upper Palaeolithic. All the large predators becomeextinct, both hypercarnivores (Panthera spelaea and P. par-dus) and meat-bone eaters as Crocuta spelaea and H. prisca.The brown bear is not reported during Magdalenian time butit has been recognized during Holocene; this is probably re-lated with the scarce faunal documentation for this time pe-

riod, recognized in very few sites (five sites, in all with tenlevels). Similar processes seem to have happened in otherregions of southern Europe (e.g., Brugal et al., 2004c), wherethe LGM peak coincides with a strong climatic threshold inthe distribution of large carnivores: reduction of range andlocal disappearance, before total extinction. However, the tim-ing of this disappearance is far from uniform throughout allWestern Europe, as demonstrated by evidences of cave hy-ena dated at c. 12 780 cal yr BP in southern Spain (LasVentanas cave, Granada; Carrion et al., 2001) or lion in Aziliandeposits dated c. 12 300 yr BP in central France (Bemilli,2000) Finally, we can also point out the difference in timing

of mammal extinctions in Portugal especially between theherbivores (preys) and the carnivores (predators) the diver-

sity diminishes at the beginning of Upper Palaeolithic for theherbivores and at the end of the same period for the carni-vores. In spite of the observed diversity pattern, it is interest-ing to note the diminishing occupation of caves by carnivoresbetween Middle and Upper Palaeolithic, thus suggesting adrastic change in term of competition of karstic places be-tween carnivorous and human groups (based on %NISP Carn/Herb: Fig. 3). On the other hand, the cavities, at least thoseoccupied both by humans or non-humans predators, are notso numerous and often have small dimensions (as expressedby the terms Lapa or Buraca which mean respectivelysmall and less deep rockshelter, and burrow or hole), or nar-row galleries. Therefore, if modern human groups still actively

seek cavities in their settlement system, complementary to theseveral open air localities, it seems that carnivores avoidsystematically such places.

In order to detail the change in mammalian associations,data have been expressed according the percentage of oc-currence, individually for herbivores and carnivores. The Ta-ble 2 indicates the most frequent taxa ranked by abundancefrom 1 to 4. The red deer is always dominant, accompaniedby horse during the Solutrean, thus indicating the coldest con-ditions for the entire last glacial. The herbivore fluctuationsdemonstrate the tendency of global climatic changes; rank-ing modifications for wild goat and aurochs, beside the asso-ciation deer/horse, are clear evidences toward gradual colderenvironment from Mousterian to Solutrean. The main differ-ence appears during the Magdalenian with more temperatespecies as boar and aurochs in detriment of horse and wildgoat, probably related to deglaciation process and climaticimprovement. With lower percentages, chamois and roe deerare also present, generally found in different levels and pro-bably more associated with local environment, namely eleva-tion and/or protected forested areas (refugium). Changes incarnivore compositions seem relatively more spectacular dueto the disappearance of several species (cf. above), especiallythe large predators often considered humans direct competi-tors. Given the data, the most interesting and logically phe-

nomenon, is the increase of lynx followed by wolf and fox, allpredators of small game like leporids. From the Early UpperPaleolithic, the trio lynx/wolf/fox becomes predominant.

Interestingly in the same time period prehistoric peoplebecome more involved too into small game acquisition andwe can enquiry about the importance of competition betweenhumans and medium to small carnivores for this type prey.This pattern can explain also the lower degree of carnivoreoccupation in caves as seen previously.

5 Capra ibexor caucasica in other parts of Europe; Capra ibexandaegagrus in Eastern Mediterranean.


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The animal communities are relatively homogenous dur-ing the Last Glacial (OIS 4-2) in Central Portugal, especially

for herbivores with persistence of temperate species as reddeer and aurochs, horse and wild goat i n the fossil record.Climatic oscillations, in relation with latitudinal and altitudinalshifts, result in modifications on the proportion of species,and subtle variations can be seen for the upper pleniglacialand late glacial. Amongst carnivores, the disappearance oflarge predators constitutes the main feature happening justafter the last cold maximum (LGM). It is important to noticethe total absence of cold adapted species as reindeer, bi-son, musk ox, saga, mammoth, woolly rhino, arctic fox, wol -verine, as well as giant deer (Megaloceros) and cave bear(Ursus spelaeus), when compared with northern region ofIberia (e.g., Cantabria). However these species were never

common in the Iberian mammal associations (except cavebear well represented in the mountain range of Pyrenees)and seem absent in the rest of the peninsula, with the nota-ble exception of a mammoth (M. primigenius ) found in a peat-bog near Granada (Padul dated to 35 790+/-960 BP, OIS 3)and woolly rhino (C. antiquitatis) from a terrace deposit nearMadrid (Arroyo Culebro, attributed to early Wrm, OIS 4?)(Garcia and Arsuaga, 2003). A clear biogeographic shiftseems to occur in Iberia peninsula during the Last Glacialwith northern region subject to colder condition (cf. the modelof Ebro frontier of some authors), more or less like the oneexisting today, which indeed reflects both climate and vege-tation zones (oceanic/eurosiberian versus mediterannean/submediterranean regions), which we can add the mountainclimate. It seems obvious that climatic conditions were rela-tively severe during al least the upper pleniglacial in Portu-gal, enough to develop grass steppes and dispersed openforest but without bringing important changes in global mam-mal associations. The southern part of Iberia was never adeserted herbivore ecosystem for the Last Glacial, but thispicture issued from many sites in Central Portugal could bea little different in other part of the countrys territory, espe-cially in northern areas.

Another determinant aspects concern specific processesinvolving large mammalian associations in Portugal. The first

one represents the survival of species in comparison withother west European faunas. The most noteworthy is the per-sistence of straight-tusked elephants (E. antiquus , Brugal andRaposo, 1999), archaic hyenids (H. prisca) and wolf (the smallform C. lupus lunellensis) from early last glacial levels of Fur-ninha (Cardoso, 1993), along with other species as rodents(A.cantiana from Mousterian levels of Figueira Brava; Jeannet,2000) or Neanderthals, until the beginning of the Last Glacial.Moreover, endemic processes are acting on species, whichpresent smallest size and slenderer elements in comparisonwith northern populations. Cervids and caprids are probablyaffected by this process and will need complementary studies,

as the one made on equids, which lead to the description ofthe new sub-species Equus caballus antunesi(Cardoso and

Eisenmann, 1989) from the early last glacial of Pedreira dasSalemas and Gruta das Fontainhas, and still recorded in alater phase at Algar de Joo Ramos (see footnote 3).

CONCLUSIONS

In the last decade, new archaeological excavations andstudies of old collections according to modern taphonomic andzooarcheological perspectives allowed new results about themammal associations supported by precise cultural layers andnumerous isotopic dates. We can then propose a comprehen-sive database about medium to large mammals covering the

Pleistocene/Paleolithic in the Portuguese territory, set upon55 sites and around 75 levels, and concerning 23 species.

The fossil record is scanty for Lower Pleistocene, and stillrare for Middle Pleistocene and the beginning of Upper Pleis-tocene. However we can point out the very important docu-mentation from the Middle Pleistocene site of Galeria Pesadawith an original association and the rich information for theperiod between late Middle Paleolithic and final Upper Pa-leolithic, with many sites located in Central Portugal.

The Last Glacial is a complex period in term of climate,well known now from high resolution data from deep-sea corelocated not far from the coast of Portugal (SU 8118, SO 75-26KL, MD 95-2042, e.g., Sanchez-Goni et al., 2000; Turon etal., 2003). Past climatic fluctuations had clear impact on landpaleoenvironment on southern regions as Iberia peninsula andAtlantic coast. We have organized our data for culturally timeperiods more or less bracketed between main cold events(Heinrich) in the last 40 Ky, thus allowing the appreciation ofdynamics in herbivore and carnivore distributions. The re-peated cold events of relatively long duration (1000-2000years) have some incidence on mammalian ecological struc-ture, at least in the variation of population density connectedwith more or less efficient rate of reproduction. To some de-gree, large mammalian association reproduce small-scale en-vironmental and climatic oscillations within the last glacial. In

Portugal, minor changes have been observed concerning thecomposition and the frequencies of species association, mar-ked by local carnivore extinctions and by a relative stabilityamong the herbivore communities without any cold-adaptedspecies known in other part of Western Europe and limited tonorthern Iberia. The main change occurs at the end of UpperPleistocene, during the development of Magdalenian culture,with the establishment of distinct mammalian populations. ThePortuguese specificity is also shown with evolutive and en-demic processes, related to biogeographical factors.

This is also a complex period regarding hominid strate-gies, as noticed in two major facts like the replacement of last


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Neanderthal groups by anatomically modern humans6 and theenlargement of diet with active acquisition of small game

during the Upper Palaeolithic, particularly the terrestrial oneas the leporids. The knowledge about past animal dynamics,related or not to climate, constitutes a relevant question inthe actual debate about settlement patterns, adaptation andfitness of human species and nature of game acquisition s.l.by prehistoric people.

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6 And probably contact, if we accept the hypothesis of crossbreeding forthe Gravettian child of Lagar Velho (Zilho and Trinkaus, 2002).


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TABLE 1. Number of occurrence of Mammal species during the Last Glacial in Portugal.

MP (n = 20) EUP (n = 15) PS + S (n = 12) Ma (n = 10) Holocene

OIS 4-(3) 4/3 2 2-(1) 1

Herbivores

Sus scrofa 7 7 5 7 +

Cervus elaphus 17 13 10 10 +

Dama dama 1* 0 0 1*

Capreolus capreolus 5 4 3 1 +

Bos primigenius 13 5 5 5 +

Capra pyrenaica 9 9 6 2-3 +

Rupicapra rupicapra 2 2 1 3

Equus caballus 14 10 10 4-5 +?

Equus hydruntinus 1 0 2 0

Stephanorhinus hemitoechus 10 1 0 0

Elephas (P.) antiquus 2 0 0 0

cf. Mammuthus (1?) 0 0 0

n. occ. sp. Herb 81 51 42 27

n. sp. Herb 11 8 8 8 5-6

Carnivores

Canis lpus 11 5 4 2 +Vulpes vulpes 10 8 5 2 +

Ursus arctos 13 3 2 0

Hyaena prisca 1 0 0 0

Crocuta spelaea 10 5 2 0

P. (Leo) spelaea 3 1 0 0

Panthera pardus 10 2 1 0

Felis silvestris 8 2 2 1 +

Lynx pardina 10 5 5 3 +

Meles meles 1 1 2 1 +

n. occ. sp. Carn 77 32 23 9

n. sp. Carn 10 9 8 5 5Rodents

Castor fiber 2 2 2 2 +

* Dama: one single remain.MP = Middle Paleolithic; EUP = Aurignacian + Gravettian; PS + S = Protosolutrean + Solutrean;Ma = Magdalenian; OIS = Oxygen Isotopic Stage.


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TABLE 2. Ranking distribution of taxa in Middle and Upper Paleolithic of Portugal(based on % occurrence per time-period).

sp. rank 1 2 3 4

Herbivores

Ma (n = 10) Cervus Sus Bos Equus

PS + S (n = 12) Cervus / Equus Capra Bos / Sus

EUP (n = 15) Cervus Equus Capra Sus

MP (n = 20) Cervus Equus Bos Capra

Carnivores

Ma (n = 10) Lynx Canis / Vulpes Felis

PS + S (n = 12) Lynx / Vulpes Canis Ursus / Crocuta / Felis

EUP (n = 15) Vulpes Canis / Lynx / Crocuta Ursus

MP (n = 20) Ursus Canis Lynx / Vulpes / Crocuta / P.pardus


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FIGURE 1. Location of Middle and Upper Palaeolithic/Pleistocene sites in Portugal (synthetic view),with line seashore at 18 Ky (after Dias et al. , 2000).1 Lorga de Dine; 2 Fujaca; 3 Mealhada; 4 Condeixa; 5 Conmbriga; 6 Buraca Escura;7 Buraca Grande; 8 Lagar Velho; 9 Gruta do Caldeiro; 10 Foz do Enxarrique; 11 Lapa

do Anecrial; 12 Lapa do Picareiro; 13 Galeria Pesada / Brecha das Lascas; 14 Gruta daNascente do Almonda; 15 Gruta da Oliveira; 16 Cabeo dos Mortos / Casais Robustos;

17 Abrigo Grande das Bocas; 18 Algar Joo Ramos; 19 Serra Molianos; 20 Furninha;21 Casa da Moura; 22 Gruta Nova da Columbeira; 23 Lapa do Suo; 24 Gruta das

Fontainhas; 25 Lapa da Rainha; 26 Quinta do Gaio; 27 Casal Torcato; 28 Carregado;29 Meirinha; 30 Santo Anto do Tojal; 31 Gruta do Pego do Diabo; 32 Gruta do

Correio-Mor; 33 Gruta das Salemas; 34 Pedreira das Salemas; 35 Algar de Cascais;36 Porto Covo; 37 Gruta da Figueira Brava; 38 Gruta do Escoural; 39 Santa Cruz;

40 Vale de Boi; 41 Algoz; 42 Goldra.


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FIGURE 2. Dated-sites ( 1 radiocarbon date, uncalibrated BP) of Middle and Upper Palaeolithic sites with faunas in Portugal(see table 1 for abbreviations); H = Heinrich events, YD = Younger Dryas.


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FIGURE 3. Mean Percentage of Carnivore versus Herbivore (based on NISP) in various cavesites of Middle and Upper Palaeolithic in Portugal and number of medium-large predators

(curve; C.lupus, U.arctos, C.spelaea, P.spelaea, P.pardus, L.pardina).


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Documents

Brugal & Valente 2007 - Dynamic of Large Mammalian Associations in the Pleistocene of Portugal